Hieracolichus Gaud and Atyeo, 1975

Mironov, Sergey V., Efeykin, Boris D., Ibanez, Jayson C., Sumaya, Anna Mae & Tolstenkov, Oleg O., 2018, Captive individuals of endangered Philippine raptors maintain native feather mites (Acariformes: Pterolichoidea) species, International Journal for Parasitology: Parasites and Wildlife 7 (2), pp. 116-133 : 118-128

Genus Hieracolichus Gaud and Atyeo, 1975

Type species: Pterolichus nisi Canestrini, 1878 , by original designation.

Representatives of the genus Hieracolichus , currently including nine species, are restricted to birds of the order Accipitriformes ( Gaud, 1983b; Gaud and Atyeo, 1974; Hernandes, 2017). Of them, seven Hieracolichus species are known from African raptors ( Gaud, 1983a). Although the genus Hieracolichus is not species-rich, taxonomic limits and species content of this genus need a revision ( Mironov et al., 2007). This genus is very close to the genus Aetacarus Gaud and Atyeo, 1975 , which has 10 of 12 known species associated with Accipitriformes . The genera Aetacarus and Hieracolichus differ from each other based only on a single feature of females: in Hieracolichus , coxal setae 4a are situated slightly anterior to the genital papillae and close to genital setae g, while in Aetacarus , these setae are situated posterior to the genital papillae, in some species even posterior to coxae IV. Because of a weak morphological boundary between two genera, Gaud (1983b) was unable to create separate keys to them and provided a single key where species of these genera were mixed together. Position of some species currently referred to the genus Hieracolichus is questionable. Thus, Hieracolichus hirundo ( Mégnin and Trouessart, 1884) placed in this genus by Gaud and Atyeo (1975) and recently redescribed by Hernandes (2017) should be formally referred to the genus Aetacarus . The redescription of this species clearly shows that in females, setae g are closer to setae 4b than 4a, and the genital papillae are situated anterior to setae 4a. These are the two main diagnostic features of Aetacarus distinguishing it from Hieracolichus . Referring of H. ostudus Gaud, 1978 to Hieracolichus , being the only species of this genus having inflated bases of epimerites I and II and lacking solenidion σ on genu III, is also doubtful.

Type material. Male holotype ( ZISP 7412), 13 male and 9 female paratypes from Pithecophaga jefferyi Ogilvie-Grant, 1896 ( Accipitridae ), THE PHILIPPINES, Agusan del Norte, Santiago, Mt. Mamajao near Lake Mainit, caught on April 1974, mite collector O.O. Tolstenkov. The bird was at least 42 years old in 2016 when the mites were sampled. Voucher specimen: paratype female ZISP 7434.

Depository: holotype, 8 male and 5 female paratypes, including voucher – ZISP, remaining paratypes – UMICHZ .

Additional material. 3 males, 1 females from P. jefferyi , THE PHILIPPINES, Lanao del Sur, Wao , wild-caught on 25 April 2015, mite collector O.O. Tolstenkov. Voucher specimen: female ZISP 7454 .

3.2. Description

MALE ( Figs. 1 and 3 A-D). (Holotype, range for nine paratypes in parentheses). Gnathosoma roughly trapezoidal, length including palps 80 (75–83), greatest width at base 78 (75–78). Idiosoma length from anterior end to bases of setae h3 on lobar apices 475 (465–490), greatest width at level of humeral setae 290 (270–290); length of hysterosoma 340 (330–350). Prodorsal shield: occupying almost entire prodorsum, Prodorsal shield: antero-lateral extensions protruding to margins of propodosoma between trochanters I and II and fused with epimerites Ia, antero-lateral margins heavily sclerotized, lateral margins with narrow and deep incisions encircling bases of setae se, posterior margin slightly sinuous, greatest length 135 (120–135), width at posterior margin 180 (170–180). Setae vi spiculiform, 70 (67–73) long, extending slightly beyond tips of palps. Setae si spiculiform, 57 (55–60) long. Distance between bases of scapular setae: se:se 87 (78–85), si:si 37 (28–35). Subhumeral setae c3 filiform, with lanceolate enlargement in basal 1/3, 100 (95–105) long. Hysteronotal shield: greatest length from anterior margins to bases of setae h3 330 (320–345), length along midline 230 (225–240), width at anterior margin 155 (150–160), anterior margin slightly concave, surface of anterior half with sparse transverse striation. Lateral bands distinct. Lobar areas of hysteronotal shield not separated from main body of hysteronotal shield. Supranal concavity small triangular. Setae c2 thin spiculiform, 70 (70–70) long, situated in anterior angles of hysteronotal shield, cupules ia immediately postero-mesal to their bases. Setae e1 situated at level of hysteronotal gland openings gl or slightly anterior to them. Length of terminal cleft from anterior end to lobar apices (setae h3) 93 (90–100), greatest width at level of setae h1 67 (65–75). Margin of anterior one third of terminal cleft heavily sclerotized, margin of remaining part membranous; this membranous margin strongly convex anterior to bases of setae h1, posterior ends of opisthosomal lobes with small semi-ovate extensions. Setae e2 spiculiform 52 (50–58) long, with apices extending slightly beyond level of setae h2; setae f2 narrowly lanceolate, 27 (27–32) long, situated at level of setae h2, setae h1 lanceolate with rounded apex, 23 (22–25) long, 3.5 (3.5–5) wide, situated posterior to level of setae h2. Distances between bases of dorsal setae and gland openings: c2:d2 120 (100–115), d2:e2 140 (140–150), e2:h3 60 (60–68), d2:gl 32 (29–35), h3:h3 95 (95–105), h2:h2 108 (100–115), d1:d2 37 (25–37), e1:e2 110 (105–115).

Epimerites I, II without inflated bases. Epimerites I with tips simple, not extending to bases of coxal setae 1a. Epimerites II slightly curved. Genital apparatus at level of trochanters IV, 23 (22–25) × 25 (25–30), aedeagus not extending to its base. Bases of setae 4a separated. Setae 4b are slightly posterior to level of setae 3a. Setae g at level of anterior pair of genital papillae. Distances between ventral setae: 4b:g 37 (35–42), g:4a 75 (67–75), 4a:ps3 37 (37–42), ps3:h3 93 (87–98), 4a: 4a 15 (13–16). Anal suckers 25 (22–25) in diameter, corolla with 18–19 rounded denticles.

Femora I, II without ventral crest. Seta cG of genu I spiculiform, 90 (85–90) long, slightly exceeding entire length of genu and tibia. Solenidion σ 1 of genu I 8 (8–11) long, much longer than solenidion σ 2. Solenidion σ of genu III situated in basal part of this segment. Solenidion φ of tibia IV s shorter than corresponding tarsus. Tarsus IV with seta d button like and seta e of minute spine-like. Legs IV with distal half of tarsus extending beyond level of lobar apices. Length of tarsi: I, II 22 (22–24), III, IV 24 (22–25). Ambulacral disc of tarsus I ovate and in longitudinal diameter noticeably longer than the more circular-shaped ambulacral discs of tarsi II–IV. Length of tarsi: I 45 (45–50), II 58 (56–59), III 62 (60–63), IV 68 (65–68). Length of

Feather mite species GenBank accession number Superfamily Family Host species

Amerodectes sp. KU202819, KU202968 Analgoidea Trouessart and Megnin, 1884 Proctophyllodidae Megnin and Vireo hypochryseus Sclater, 1863

Trouessart, 1884

Amerodectes turdinus (Berla, 1959) KU 203310 Analgoidea Trouessart and Megnin, 1884 Proctophyllodidae Megnin and Catharus fuscescens Stephens, 1817

Trouessart, 1884

Ascouracarus sp. JQ000778, JQ000475, JQ000167 Pterolichoidea Trouessart and Mégnin, 1884 Ascouracaridae Gaud and Atyeo, 1976 Strix virgata Cassin, 1850

Cystoidosoma sp. JQ000777, JQ000474, JQ000166 Pterolichoidea Trouessart and Mégnin, 1884 Ascouracaridae Gaud and Atyeo, 1976 Melanerpes aurifrons Wagler, 1829

Mesosathes sp. JQ000753, JQ000448 Pterolichoidea Trouessart and Megnin, 1884 Crypturoptidae Gaud, Atyeo and Berla , Crypturellus boucardi Sclater, 1860

1972

Falculifer sp. JQ000748, JQ000135, JQ000135 Pterolichoidea Trouessart and Mégnin, 1884 Falculiferidae Oudemans, 1908 Columba flavirostris Wagler, 1831

Falculifer sp. JQ000749 Pterolichoidea Trouessart and Mégnin, 1884 Falculiferidae Oudemans, 1908 Columba oenas Linnaeus, 1758

Falculiferidae sp. JQ000751, JQ000138 Pterolichoidea Trouessart and Mégnin, 1884 Falculiferidae Oudemans, 1908 Scardafella inca Lesson, 1847

Hyperaspidacarus sp. JQ000750, JQ000137, JQ000445 Pterolichoidea Trouessart and Mégnin, 1884 Falculiferidae Oudemans, 1908 Scardafella inca

Freyana anatina (Koch, 1844) JQ 000743, JQ000438 Pterolichoidea Trouessart and Mégnin, 1884 Freyanidae Dubinin, 1951 Anas platyrhynchos Linnaeus, 1758

Freyana lophodytes Dubinin, 1950 JQ 000746, JQ000441, JQ000133 Pterolichoidea Trouessart and Mégnin, 1884 Freyanidae Dubinin, 1951 Lophodytes cucullatus Linnaeus,

1758

Freyana sp. JQ000744, JQ000439 Pterolichoidea Trouessart and Mégnin, 1884 Freyanidae Dubinin, 1951 Aix sponsa Linnaeus, 1758

Freyana sp. JQ000442 Pterolichoidea Trouessart and Mégnin, 1884 Freyanidae Dubinin, 1951 Tadorna ferruginea Pallas, 1764

Freyanidae Dubinin, 1951

Aetacarus sp. JQ000769, EU152516, JQ000465 Pterolichoidea Trouessart and Mégnin, 1884 Gabuciniidae Gaud and Atyeo, 1975 Geranospiza caerulescens Vieillot ,

1817

Capitolichus sp. JQ000774, JQ000470 Pterolichoidea Trouessart and Mégnin, 1884 Gabuciniidae Gaud and Atyeo, 1975 Dryocopus lineatus Linnaeus, 1766

Capitolichus sp. JQ000161 Pterolichoidea Trouessart and Mégnin, 1884 Gabuciniidae Gaud and Atyeo, 1975 Melanerpes aurifrons Wagler, 1829

Coraciacarus americanu s Alzuet, Cicchino and EU152770, JQ000165, JQ000473 Pterolichoidea Trouessart and Mégnin, 1884 Gabuciniidae Gaud and Atyeo, 1975 Coccyzus americanus Linnaeus ,

Abrahamovich, 1988 1758

Gabucinia delibata (Robin, 1877) JQ 000770, JQ000158, JQ000466 Pterolichoidea Trouessart and Mégnin, 1884 Gabuciniidae Gaud and Atyeo, 1975 Corvus brachyrhynchos Brehm ,

1822

Gabucinia sp. JQ000771 Pterolichoidea Trouessart and Mégnin, 1884 Gabuciniidae Gaud and Atyeo, 1975 Cyanocorax sanblasianus

Lafresnaye, 1842

Hieracolichus nisi (Canestrini, 1878) JQ 000776, JQ000164, JQ000472 Pterolichoidea Trouessart and Mégnin, 1884 Gabuciniidae Gaud and Atyeo, 1975 Accipiter nisus Linnaeus, 1758

Hieracolichus philippinensis MF 967008, MF967010, MG001908, MG001910, Pterolichoidea Trouessart and Mégnin, 1884 Gabuciniidae Gaud and Atyeo, 1975 Pithecophaga jefferyi

MG001915, MG001917

Piciformobia sp. JQ000775, JQ000163, JQ000471 Pterolichoidea Trouessart and Mégnin, 1884 Gabuciniidae Gaud and Atyeo, 1975 Crotophaga sulcirostris Swainson ,

1827

Dermonoton sp. JQ000742, JQ000437, JQ000129 Pterolichoidea Trouessart and Mégnin, 1884 Kramerellidae Gaud and Mouchet, 1961 Glaucidium brasilianum Gmelin ,

1788

Kramerella oti (Lönnfors, 1937) JQ 000740, JQ000435 Pterolichoidea Trouessart and Mégnin, 1884 Kramerellidae Gaud and Mouchet, 1961 Asio otus Linnaeus, 1758

Kramerella sp. JQ000128, JQ000436 Pterolichoidea Trouessart and Mégnin, 1884 Kramerellidae Gaud and Mouchet, 1961 Bubo virginianus Gmelin, 1788

Pseudogabucinia nisaeti MF 967012, MG001912, MG001919 Pterolichoidea Trouessart and Mégnin, 1884 Kramerellidae Gaud and Mouchet, 1961 Nisaetus pinskeri

Geranolichus canadensis Atyeo and Windingstad, JQ 000755, JQ000142, JQ0004501 Pterolichoidea Trouessart and Megnin, 1884 Pterolichidae Trouessart and Megnin , Grus canadensis Linnaeus, 1758

1979 1884

Grallobia fulicae (Trouessart, 1885) JQ 000757 Pterolichoidea Trouessart and Megnin, 1884 Pterolichidae Trouessart and Megnin , Fulica atra Linnaeus, 1758

1884

Grallobia sp. JQ000756 Pterolichoidea Trouessart and Megnin, 1884 Pterolichidae Trouessart and Megnin , Porzana carolina Linnaeus, 1758

1884

Grallolichus sp. JQ000758, JQ000145, JQ000453 Pterolichoidea Trouessart and Megnin, 1884 Pterolichidae Trouessart and Megnin , Gallinula chloropus Linnaeus, 1758

1884

Kakapolichus sp. JQ000759, JQ000454 Pterolichoidea Trouessart and Megnin, 1884 Pterolichidae Trouessart and Megnin , Nestor notabilis Gould, 1856

1884

Pseudalloptinus pithecophagae MF 967007, MF967009, MF9670011, MF9670013, Pterolichoidea Trouessart and Megnin, 1884 Pterolichidae Trouessart and Megnin , Pithecophaga jefferyi

MG001920, MG001914, MG001909, MG001918 1884

Pterolichus obtusus Robin, 1877 JQ 000754, EU152513, JQ000449 Pterolichoidea Trouessart and Megnin, 1884 Pterolichidae Trouessart and Megnin , Gallus gallus Linnaeus, 1758

1884

Aniacarus mexicanus Gaud and Atyeo, 1990 JQ 000762, JQ000457 Pterolichoidea Trouessart and Mégnin, 1884 Pterolichidae Trouessart and Mégnin , Crotophaga sulcirostris Swainson ,

1884 1827

(continued on next page)

, 1758

Linnaeus, 1758,

1758 1758, Linnaeus Linnaeus,

sulcirostris autumnalis Herodias Linnaeus autumnalis sulcirostris auritus Linnaeus Linnaeus, squatarola vociferus minuta Leisler, 1812, totanus 1758 Linnaeus Host species Crotophaga Amazona 1758 Ardea Amazona Crotophaga atra Fulica Podiceps Aix sponsa Pluvialis Charadrius 1758 Calidris Tringa Charadrius vociferus

,,

,

Mégnin Mégnin

,

Mégnin

,

Mégnin

, Mégnin Mégnin and and and and and and 1896,, 1896 1896,, 1896, 1896 Trouessart Trouessart Trouessart Trouessart Trouessart Trouessart 1982, Gaud 1961 Gaud, Trouessart Trouessart Trouessart Trouessart Trouessart Family Pterolichidae 1884 Pterolichidae 1884 Pterolichidae 1884 Pterolichidae 1884 Pterolichoidae 1884 Pterolichoidae 1884 Ptiloxenidae Rectijanuidae Syringobiidae Syringobiidae Syringobiidae Syringobiidae Syringobiidae 1884 1884 1884 1884 1884 1884 1884 1884 1884 1884 1884 1884 1884,,,,,,,,,,,,, Mégnin Mégnin Mégnin Mégnin Mégnin Mégnin Megnin Megnin Megnin Megnin Megnin Mégnin Mégnin and and and and and and and and and and and and and Trouessart Trouessart Trouessart Trouessart Trouessart Trouessart Trouessart Trouessart Trouessart Trouessart Trouessart Trouessart Trouessart Superfamily Pterolichoidea Pterolichoidea Pterolichoidea Pterolichoidea Pterolichoidea Pterolichoidea Pterolichoidea Pterolichoidea Pterolichoidea Pterolichoidea Pterolichoidea Pterolichoidea Pterolichoidea

000148 000139 000147 000152 000463 000156 000153 number, JQ, JQ JQ, JQ, JQ,, JQ JQ, accession 000456 JQ, 000447 JQ,, JQ 000455 000460, JQ, 000459 JQ, JQ 000155 000464, JQ, JQ 000154, 000461 JQ GenBank 000763 JQ 000761 JQ JQ 000752 JQ 000760 000458 JQ JQ 000144 000764 JQ EU152767 000767 JQ 000768 JQ 000766 JQ 000765 JQ JQ 000464 1990 1990 1993

,

, Atyeo, Atyeo) and Atyeo 1904

and and, 1885 Megnin and, Oudemans

)

Gaud

sp.

Gaud

Trouessart longitarsa

(

) 1884

Dabert

Table

3

continued

(

mite

Feather species

Aniibius drepanophorus Chelomatolichus sp.

Herodialges Scolaralichus

sp.

Aniibius drepanophorus fulicae

Grallobia

(sp

Ptiloxenus

. sp.

Rectijanua Leptosyringobia

Trouessart,

Phyllochaeta tenuiseta .

Syringobiidae

sp

Plutarchusia chelopus

Syringobiidae sp.

solenidia: σ 1 I 18 (13–18), σ II 6 (5–7), σ III 20 (15–20), ω 1 I 12 (11–14), ω 1 II 25 (24–26).

FEMALE ( Figs. 2 and 3E, F ) (range for nine paratypes). Gnathosoma, length × width, 105–110 × 100–102. Idiosoma, length × width, 615–640 × 340–355, length of hysterosoma 430–445. Prodorsal shield shaped as in male, 155–165 × 185–200. Setae vi spiculiform, 75–80 long, barely reaching tips of palps. Setae si spiculiform, 75–78 long. Distance between bases of scapular setae: se:se 105–120, si:si 35–38. Subhumeral setae c3 filiform, 115–125 long. Hysteronotal shield: main body with almost straight anterior margin, anterior angles acute, posterior end extending to midlevel between hysteronotal gland openings gl and setae e2, posterior margin with blunt-angular median extension and pair of shallow concavities, greatest length 360–370, width at anterior margin 270–280, surface with faint transverse striation. Setae c2 spiculiform, 92–105 long, situated off hysteronotal shield; cupules ia postero-mesal to them and also off this shield. Setae d2 short filiform, about 20 long. Setae e1 approximately at level of hysteronotal gland openings gl. Lateral bands well developed, longer than main body of hysteronotal shield, with posterior ends almost extending to cupules ip and slightly curved medially. Posterior one quarter of opisthosoma poorly sclerotized, with fine striation and, in some specimens, with barely distinct punctation. Setae e2 spiculiform, 115–125 long, setae f2 filiform 30–40 long, setae h1 short filiform, about 10 long; both pair situated on poorly sclerotized area of opisthosoma. Posterior end of opisthosoma with wide and rounded median extension bearing setae h2, h3 and ps1 and with strongly sclerotized margin. Distances between dorsal setae and gland openings: c2:d2 135–155, d2:e2 155–170, e2:h3 78–83, d2:gl 72–78, h1:h1 62–70, h2:h2 75–80. h3:h3 45–48.

Epimerites I, II without basal inflation. Epimerites I not extending to setae 1a. Epigynum horseshoe-shaped, 72–88 long, 92–100 wide. Setae 4b situated on epigynum, close to its tips. Setae 4a situated slightly anterior to genital papillae. Copulatory opening immediately posterior to anal opening. Distances between ventral setae: 4b:g 75–80, 4b:3a 35–50, g:4a 13–25, ps2:ps3 27–32, ps2:ps2 67–72.

Femora I, II with ventral crest. Setae cG of long spiculiform, 22–28 long, approximately subequal to entire length of genu and tibia I. Legs IV with tarsus and distal part of tibia extending beyond posterior end of opisthosoma. Length of tarsi: I 53–58, II 72–78, III 75–80, IV 93–100. Length of solenidia: σ 1 I 23–28, σ II 8–12, σ III 17–25, ω 1 I 16–18, ω 1 II 22–24.

Differential diagnosis. Among previously described species, Hieracolichus philippinensis sp. n. is more similar to H. dobyi Gaud and Mouchet, 1959 described from Stephanoaetus coronatus (Linnaeus, 1766) in Africa ( Gaud and Mouchet, 1959; Gaud, 1983b) in having, in males, setae e2 extending to the level of setae h2 and f2, and relatively short and narrowly lanceolate setae h1. Hieracolichus philippinensis differs from this species by the following features: in both sexes, setae c3 are long, filiform and exceed 100 μm in length, and genual solenidion σ is situated at the base of genu III; in males, setae g are situated almost at the level of anterior genital papillae; setae h1 are short (22–25 μm), and the inner margins of opisthosomal lobes have a pair of noticeably convex membranes in the anterior part of the terminal cleft; in females, the hysteronotal shield is shaped as an inverted trapezium and the posterior one third of the opisthosoma is devoid of sclerotization except the posterior margin, and tarsus IV completely extends beyond the posterior margin of the opisthosoma. In both sexes of H. dobyi , setae c3 are narrowly lanceolate at base with filiform apex (80-90 μm long), and genual solenidion σ is situated at the midlength of genu III; in males, setae g are situated anterior to the level of genital papillae; setae h1 are narrowly lanceolate, curved and 30–35 μm long, and the inner margins of opisthosomal lobes are almost straight; in females, the hysteronotal shield is shaped as an inverted trapezium and the posterior one third of the opisthosoma is devoid of sclerotization except for the posterior margin, and tarsus IV slightly (by ¼ the length) extends beyond the posterior margin of the opisthosoma.

Etymology. The specific epithet is derived from the country, where the mite was found.

Family Pterolichidae Trouessart et Mégnin, 1884

Subfamily Pterolichinae Trouessart et Mégnin, 1884

Genus Pseudalloptinus Dubinin, 1956

Type species: Pterolichus (Pseudalloptes) aquilinus var. milvulinus Trouessart, 1884 , by original designation.

The genus Pseudalloptinus originally included pterolichine mites associated with birds from the orders Accipitriformes , Falconiformes , Gruiformes , Ciconiiformes and Psittaciformes ( Dubinin, 1956; Gaud and Mouchet, 1959). After a revision ( Gaud, 1988), the content of this genus was reduced to five species associated exclusively with birds of the order Accipitriformes . The genus Pseudalloptinus is readily distinguishable from other pterolichine genera in having, in most species, a unique structure in males: the postgenital sclerite [= fossette post-genitale of Gaud (1988)]. This sclerite, being apparently a derivative of adanal apodemes, is situated between the genital apparatus and anal field and usually is stirrup-shaped or roughly ovate.

Type material. Male holotype (ZISP 7330), 20 male and 20 female paratypes from Pithecophaga jefferyi Ogilvie-Grant, 1896 ( Accipitridae ), THE PHILIPPINES, Lanao del Sur, Wao, 25 April 2015, mite collector O.O. Tolstenkov. Voucher specimen: female paratype ZISP 7371.

Depository. Holotype, 15 male and 15 female paratypes – ZISP, remaining paratypes – UMICHZ.

Additional material. 20 males, 20 females from 3 P. jefferyi individuals originated from the following locations: 10 males, 10 females – THE PHILIPPINES, Agusan del Norte, Santiago, Mt. Mamajao near Lake Mainit , caught on April 1974 ; 5 males, 5 females, THE PHILIPPINES, Davao Oriental, Mati, Don Salvador, South Biasong , caught on 13 January 2011 ; 5 males, 5 females, THE PHILIPPINES, Davao City, Malagos, Philippine Eagle Center , 4 February 2002 (captive breed) , mite collector O.O. Tolstenkov. Voucher specimens: female paratypes ZISP 7391 , 7411 .

MALE ( Fig. 4 , 6A–C ). (Holotype, range for eight paratypes in parentheses). Gnathosoma: length including palps 62 (60–65), greatest width at base 47 (46–50). Idiosoma length from anterior end to lobar apices (bases of setae h3) 25 (325–350), greatest width at level of humeral setae 180 (180–195). Length of hysterosoma 215 (210–225). Prodorsal shield: occupying most part of prodorsum, antero-lateral extensions acute, lateral margins with deep and narrow extensions encircling bases of scapular setae se, posterior margin slightly concave, length along midline 98 (95–105), greatest width 102 (100–110). Setae vi filiform, 38 (28–38) long, not extending to palpal apices. Setae se separated by 65 (65–68). Setae si minute filiform, close to bases of corresponding setae se. Scapular and humeral shields present. Setae c2 filiform, 15 (12–15) long, situated on anterior margin of humeral shields. Subhumeral setae c3 lnceolate 20 (18–20) long, 4 (3.7–5) wide.

Hysteronotal shield: greatest length from anterior margins to bases of setae h3 212 (210–215), width at anterior margin 145 (140–150), anterior margin slightly concave, surface without ornamentation. Lateral bands distinct, narrow. Hysteronotal gland opening gl situated at level of trochanters IV. Setae d2 minute filiform about 10 (10–12) long; setae e2 filiform, 16 (15–18) long, situated at level of anterior end of supranal concavity, not extending to lobar apices. Opisthosomal lobes roughly triangular, at base slightly wider than long, rounded posteriorly. Terminal cleft roughly semi-ovate, 28 (25–30) long, 35 (34–38) in width at level of setae ps1. Supranal concavity open posteriorly into terminal cleft. Terminal cleft with narrow entire membrane forming semi-ovate terminal extensions on lobar apices, length of these extensions 10 (10–15) long, wide at base 18 (17–20). Setae ps2 long filiform, extending far beyond level of lobar apices; setae ps1 minute filiform, about 15 long, situated near bases of setae h2. Distances between dorsal setae: c2:d2 87 (80–88), d2:e2 75 (72–80), e2:h3 42 (40–45), ps1:ps1 40 (38–42), h2:h2 60 (60–65), h3:h3 50 (50–55), ps2:ps2 70 (70–75).

Epimerites I fused into a Y with short stem. Epimerites IIa present. Genital apparatus situated at level of anterior margin of trochanters IV, 14 (14–15) long, 14 (13–17) wide. Setae 4b slightly posterior to level of setae 3a. Setae g equidistant from genital arch apex and level of setae 4b. Anterior genital papillae at level of genital arch apex. Epimerites IVa long, bearing bases of setae 4a near tips and flanking base of genital arch. Adanal apodemes with L-shaped inner ends flanking median area with bases of setae ps3 but not forming separate postegenital sclerite. Anal suckers 13 (13–15) in diameter, corolla without indentation, surrounding membrane very wide and extending laterally over lateral margins of opisthosoma. Distances between ventral setae: 4b:g 23 (22–25), g:4a 40 (40–47); 4a:ps3 30 (30–32), ps3:h3 67 (65–68).

Setae of tarsi I, II filiform. Solenidion σ 1 situated at its midlevel of genu I and 1.3–1.5 times longer than this segment. Genual setae cG I, cG II, mG I and mG II filiform, shorter than corresponding segments. Solenidion σ of genu III in distal part of segment. Legs IV with distal half of tarsus extending beyond level of lobar apices. Tarsus IV with clawlike apical extension, setae d and e minute are absent. Solenidion φ of tibia IV about 1.5 times longer than tarsus IV. Length of tarsi: I 35 (35–37), II 35 (35–38), III 38 (37–40), IV 33 (32–34). Length of solenidia: σ 1 I 40 (40–45), σ II 8 (7.5–8), σ III 8 (8–10), ω 1 I 11 (10–03), ω 1 II 18 (16–18).

FEMALE ( Figs. 5 and 6 G,H ). Gnathosoma, length × width, 82–85 × 67–72. Idiosoma, length × width, 510 × 550. Length of hysterosoma 325–365. Prodorsal shield: shaped as in male, but lateral margins without deep incisions, 135–145 long, 135–140 wide. Setae se separated by 80–85; setae si minute filiform, situated closely to corresponding setae se. Scapular and humeral shields present. Setae c2 short filiform, 20 (18–20) long, situated in anterior margin of humeral shields. Subhumeral setae c3 lanceolate, 26–30 long, about half the length of humeral setae cp. Hysteronotal shield: entire, extending to posterior end of opisthosoma, anterior margin concave, 300–340 long, 210–220 wide at anterior margin surface without ornamentation, posterior end with desclerotized transverse area bearing setae e2. Setae c1 on hysteronotal shield near its anterior margin. Setae d2 situated approximately at midlevel between cupules ip and hysteronotal gland openings gl. Setae e2 filiform, about 10–12 long. Lateral bands present, poorly distinct. Posterior margin of opisthosoma with relatively wide terminal extension bearing setae h2, h3 and ps1. External copulatory tube minute, situated terminally about 2–3 long. Spermatheca and spermaducts as in Fig. 6H , length of secondary spermaducts 10–12. Length of opisthosomal setae: e2 18 –20, f2 8–10, ps1 5–6, ps2 15–18. Distances between dorsal setae and openings: c2:d2 175–190, d2:e2 80–95, d2:gl 34–36, h2:h3 40–52, h2:h2 35–38, h3:h3 17–18.

Epimerites I as in male. Epimerites IVa present. Epigynum semicircular, thin, 42–48 long, 65–80 wide, with tips extending to level of setae 4b. Apodemes of oviporus narrow, barely sclerotized. Setae g situated approximately equidistant from levels of setae 4b and g. Distances between ventral setae: 4b:g 50–58, 4b:3a 52–65, g:4a 38–52.

Legs I–III as in male. Solenidion σ of genu III in distal part of segment. Solenidion φ of tibia III slightly longer than corresponding tarsus; solenidion φ of tibia IV about 1/5 the corresponding tarsus. Legs IV with tarsus and distal half of tibia extending beyond posterior end of opisthosoma. Legs I, as in male. Length of tarsi: I 50–53, II 50–55, III 57–60, IV 78–80. Length of solenidia: σ 1 I 68–80, σ II 10–15, σ III 10–18, ω 1 I 12-14, ω 1 II 26–28.

Differential diagnosis. The new species, Pseudalloptinus pithecophagae sp. n. is most similar to P. africanus Gaud, 1988 and P. milvulinus (Trouessart,1884) in having the following features: in both sexes, setae c3 are lanceolate; in males, opisthosomal lobes are well developed, with semi-ovate terminal membranes; and in females, the striated sejugal area is large and constitutes about 1/5th of the total length of the idiosoma. Pseudalloptinus pithecophagae sp. n. differs from these species by the following features: in males, the genital apparatus is situated at the level of the anterior margin of trochanters IV, epimerites IVa are long and almost extending to the genital arch, and setae e2 are filiform, situated at the level of the anterior end of supranal concavity and not do not extend to lobar apices; in females, the hysteronotal shield is entire, the epigynum is semicircular and extends to the level of setae 4b, setae c1 is situated on the hysteronotal shield, external copulatory tube is minute (only 2–3 μm long), and setae g are situated at the level of setae 3a. In males of P. africanus and P. milvulinus , the genital apparatus is situated at the level of the posterior margin of trochanters III, epimerites IVa are poorly developed, and setae e2 are spiculiform, situated posterior to the supranal concavity and extend beyond the lobar apices; in females, the hysteronotal shield is spit into a large anterior piece and a small pygidial fragment covering the very posterior end of the opisthosoma, the epigynum is bow-shaped and does not extend to the level of setae 4b, setae c1 are situated on striated tegument near the anterior margin of the hysteronotal shield, the external copulatory tube is about 15 μm long and curved ventrally, and setae g are situated posterior to the level of setae 3a.

The unique feature of P. pithecophagae males, easily discriminating this species from all previously known Pseudalloptinus species, is the absence of the entire postgenital sclerite well separated from the adanal apodemes. In this species, L-shaped tips of adanal apodemes turned anteriorly and flank small median area with setae ps3, apparently corresponding to the lateral pieces of the postgenital sclerite of other species of this genus.

Etymology. The specific epithet is derived from the generic name of the type host and is a noun in the genitive case.

Family Kramerellidae Gaud et Mouchet, 1961

Genus Pseudogabucinia Č erny, 1961

Type species: Pterolichus ciconiae Canestrini et Berlese, 1881 , by monotypy.

Up to now, the feather mite genus Pseudogabucinia has included only five species with hosts erratically distributed among non-passerine orders: Accipitriformes , Ciconiiformes , Falconiformes , Gruiformes , and Otidiformes ( Table 4) ( Atyeo and Windingstad, 1979; Canestrini and Berlese, 1881; Dubinin, 1956; Gaud, 1968, 1983a; Gaud and Mouchet, 1961; Mégnin and Trouessart, 1884). This type of distribution is in surprising contrast to other six genera of Kramerellidae , each of which is associated with a particular bird order ( Gaud and Atyeo, 1996).

Among previously known Pseudogabucinia species, Pseudogabucinia intermedia ( Mégnin et Trouessart, 1884) has been recorded from raptor birds of two orders: from falcons Falco ( Falconiformes : Falconidae ), harriers Circus ( Accipitriformes : Accipitridae ) and buzzards Buteo ( Gaud, 1988) . Association of one species on hosts from different orders is quite rare among feather mites; therefore, it cannot be excluded that P. intermedia from these hosts ( Table 4) could represent separate species. In the differential diagnosis below, the new species is compared with the specimens of P. intermedia from falcons.

Type material. Male holotype (ZISP 7307), 4 male and 1 female paratypes from Nisaetus pinskeri (Gould, 1863) . ( Accipitridae ), THE PHILIPPINES, Salaysay, Davao City, caught in 2005, mite collector O.O. Tolstenkov. Voucher specimen: female paratype ZISP 7312.

Depository. Holotype, 3 male and 1 female paratypes – ZISP, 1 male paratype UMICHZ.

MALE ( Fig. 7 , 9 A-E). (Holotype, range for three paratypes in parentheses). Gnathosoma: length including palps 43 (42–45), greatest width at base 50 (48–52). Idiosoma length from anterior end to lobar apices (bases of setae h3) 270 (265–280), greatest width at level of humeral setae 175 (170–180); length of hysterosoma 195 (190–195). Prodorsal shield: occupying anterior part of prodorsum, roughly trapezoidal in shape, with slightly convex posterior margin and posterior angles slightly extending laterally, not extending to bases of scapular setae, length along midline 45 (45–48), greatest width 47 (45–50) ( Fig. 7 ). Setae se separated by 57 (55–58). Setae si spiculiform, 35 (35–47) long, separated by 23 (22–25), approximately equidistant from midline and corresponding setae se. Scapular and humeral shields absent. Setae c2 spiculiform, 30 (27–32) long, situated in striated tegument. Subhumeral setae long filiform, nearly half the length of macrosetae cp. Hysteronotal shield: greatest length from anterior margins to bases of setae h3 185 (180–190), width at anterior margin 125 (115–125), anterior margin slightly concave, lateral margins almost straight, surface with fine longitudinal striae between levels of setae e1 and e2. Supranal concavity narrowed anteriorly and extending to level of setae e1. Hysteronotal gland opening gl situated approximately equidistant from levels of setae d2 and e2. Lateral bands poorly demarcated. Seta d2 minute filiform, about 5 long, setae e2 filiform 32 (27–33). Opisthosomal lobes roughly triangular, with rounded posterior ends, approximately as long as wide at base; apical and inner margins of lobes membranous. Terminal cleft wide triangular, with blunt anterior very end, 52 (52–55) long, 52 (50–55) in width at level of setae h3. Setae f2 narrowly lanceolate with short filiform apex 40 (32–40); setae ps2 blade-shaped 27 (25–27); setae h1 narrowly triangular, 15 (15–18) long thin, setae ps1 filiform, about 10 long, situated posterior to level of setae h1. Distances between dorsal setae: c2:d2 77 (70–80), d2: e2 57 (55–60), e2:h3 63 (57–63), d1:d2 37 (35–40), e1: e2 25 (22–28), f2:f2 112 (110–120), ps1:ps1 85 (82–88), h3:h3 72 (70–75), h2:h2 105 (100–105).

Epimerites I free, slightly converging. Epimerites IIa present, barely distinct. Genital apparatus 15 (14–15) in length, 13 (13–17) in width, its base situated at midlevels of trochanters IV ( Fig. 7B ). Setae 3a and 4b situated at the same level. Setae g at level of apex of genital arch. Genital papillae situated lateral to anterior half of genital arch. Distances between ventral setae: 4b:g 17 (16–18), g:4a 23 (20–23); 4a:ps3 62 (60–64), ps3:h3 47 (47–50). Anal suckers 13 (12–14) in diameter, corolla with two rounded denticles. Small adanal sclerites presents between setae ps3 and anal suckers.

Solenidion σ 1 of genu I approximately half the length of this segment. Setae mG of genu II much longer than of genu I. Setae cG of genua I and III filiform, slightly longer than corresponding segments. Solenidion φ of tibia IV slightly shorter than tarsus IV. Setae d and e of tarsi IV minute spine-like. Legs IV with ambulacral disc slightly extending beyond level of lobar apices. Length of tarsi: I 33 (32–34), II 42 (40–43), III 40 (37–40), IV 43 (40–43). Length of solenidia: σ 1 I 5 (5–6), σ II4 (4–6), σ III 5 (5–6), ω 1 I 12 (12–14), ω 1 II 20 (18–20).

FEMALE ( Fig. 8 ). Gnathosoma, length × width, 55 × 63. Idiosoma, length × width, 310 × 200, length of hysterosoma 230. Prodorsal shield: shaped as in male, 55 × 58. Setae se separated by 68; setae si spiculiform, 45 long, separated by 30, situated approximately equidistant from midline and corresponding setae se. Scapular and humeral shields absent. Setae c2 thin spiculiform, 35 long, situated in anterior angles of humeral shields. Subhumeral setae c3 long filiform 37 long, about half the length of setae cp. Hysteronotal shield: length 180, width 125, anterior margin nearly straight, not extending to level of setae c2, surface without ornamentation, posterior margin with pair of narrow incision almost extending to level of setae e1 and wide semi-rounded extension between them. Setae d2 off hysteronotal shield. Lateral bands present, poorly demarcated. Spermatheca and spermaducts as in Fig. 9H , secondary spermaducts heavily sclerotized. Length of opisthosomal setae: e2 38, f2 125, ps1 40, ps2 155, h1 10. Distances between dorsal setae: c2:d2 87, d2:e2 83, e1: e2 20, h1:h1 50, h2:h2 83, h3:h3 55, ps1:ps1 32.

Epimerites I as in male. Epigynum bow-shaped, situated between tips of epimerites II, 15 long, 40 wide. Apodemes of oviporus barely sclerotized. Setae g and 3a situated approximately at same level of setae. Distances between ventral setae: 4b:g 12, 4b:3a 15, g:4a 32. Legs I–III as in male. Solenidion φ of tibia III slightly longer than corresponding tarsus; solenidion φ of tibia IV about one third the corresponding tarsus. Legs IV with ambulacral disc extending beyond posterior end of the opisthosoma.

Length of tarsi: I 35, II 50, III 45, IV 50. Length of solenidia: σ 1 I 10, σ II 9, σ III 6, ω 1 I 12, ω 1 II 20.

Differential diagnosis. The new species, Pseudogabucinia nisaeti sp. n. is close to P. intermedia ( Mégnin et Trouessart, 1884) known from falcons by in having, in both sexes, ambulacral discs of tarsi IV extending to or slightly beyond the posterior margin of the body, and setae c2 exceeding the distance between internal scapular setae si, and, in females, setae f2 and ps2 being equal to or exceeding the distance between their bases. Pseudogabucinia nisaeti sp. n. differs from that species by the following features: in both sexes, subhumeral setae c3 are long filiform and approximately half as long and humeral setae cp, solenidion ω 1 of tarsus II does not extend to the apex of this segment; in males, the supranal concavity does not extend beyond the level of setae e1, setae 4a are situated posterior to the base of the genital arch; in females, the genital papillae are situated distinctly anterior to the level of setae g. In both sexes of P. intermedia , subhumeral setae c3 are about 1/3 the length of setae cp, solenidion ω1 of tarsus II extends to the apex of this segment; in males, the supranal concavity extend far beyond the level of setae e1, setae 4a are situated posterior at the of the level of genital arch base; in females, the genital papillae are situated at the level of setae g.

Etymology. The specific epithet is derived from the generic name of the type host and is a noun in the genitive case.

Figures

Fig. 1. Hieracolichus philippinensis sp. n. male. A – dorsal view, B – ventral view.

Fig. 3. Hieracolichus philippinensis sp. n. details. A – opisthosoma of male, dorsal view B–D – genua, tibiae and tarsi I–III of male, respectively, E – tibia and tarsus IV of male, G – tibia and tarsus IV of female, H – spermatheca and spermaducts.

Fig. 2. Hieracolichus philippinensis sp. n. female. A – dorsal view, B – ventral view.

Fig. 4. Pseudalloptinus pithecophagae sp. n. male. A – dorsal view, B – ventral view.

Fig. 6. Pseudalloptinus pithecophagae sp. n. details. A – opisthosoma of male, ventral view, B–D – legs I–III of male, respectively, E – tibia and tarsus IV of male, G – tibia and tarsus IV of female, H – spermatheca and spermaducts.

Fig. 5. Pseudalloptinus pithecophagae sp. n. female. A – dorsal view, B – ventral view.

Fig. 7. Pseudogabucinia nisaeti sp. n. male. A – dorsal view, B – ventral view.

Fig. 9. Pseudogabucinia nisaeti sp. n. details. A – opisthosoma of male, ventral view, B–D – genu, tibia and tarsus I–III of male, respectively, dorsal view, E – tibia and tarsus IV of male, F, G – tibia and tarsus III and IV of female, respectively, G – tibia and tarsus IV of female, H – spermatheca and spermaducts.

Fig. 8. Pseudogabucinia nisaeti sp. n. female. A – dorsal view, B – ventral view.